An Overview of Late Cretaceous and Early Palaeogene Echinoderm Faunas from Liege-Limburg (Belgium, the Netherlands)

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BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE SCIENCES DE LA TERRE, 69-SUPP. A: 103-118, 1999 BULLETIN VAN HIT KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN AARDWETENSCHAPPEN. 69-SUPP. A: 103-1 IX. 19»

An overview of Late Cretaceous and Early Palaeogene echinoderm faunas from Liege-Limburg (Belgium, The Netherlands)

by John W. M. JAGT

Abstract My3eHHHM KOJUieKIJKHM, H B OC06eHHOCTH C034aHHbIM RO 1975 roaa, He XBaraeT, B nacTHOCTH, noapoôHoft HHthopMauHH o With the exception of echinoids, echinoderm faunas from the type area CTpaTHrpatnHHecKOM npoHcxo»c;reHHH. HoBaa KOJLieKima He of the Maastrichtian Stage still are more or less terra incognita. TOJibKO SHaHHTeiiBHO \TJiy6jiHeT Hanm 3H3HH5I O tbavHax Material collected recently in the area by a group of professional and HraoK05KHX LIo3ÄHero Mena (KaMnaHCKo-MacipHXTCKHH apycbi) amateur palaeontologists comprises numerous new records, which H Parmero riarteoreHa (TJaTCKMH apyc) B aaHHOH oônactn, HO H have the added advantaue of being well documented stratigraphically. Museum collections, and those pre-dating 1975 in particular, generally no3BOjmeT nojp3ecTH HTOTH no cTpyKTvpe pa3Hoo6pa3H« H suffer from a lack of detail where slratigraphic provenance is con• BbiMHpaHH», npejiiiiecTBOBaBuieH rpamme K/T H BKpecT rparame cerned. Not only do these new collections considerably increase our K/T. Kpancoe o6o3peiöie dpavH HTJIOKOJKHX npe^cTaBaeHO B knowledge of Late Cretaceous (Campanian-Maastrichtian) and Early aaHHOM OMepKe, ocoôoe BHHMaHHe yaeaeHO MopcKHM e*aM H Palaeogene (Danian) echinoderm faunas in the area, they also allow acTepoH/raM. conclusions on diversification and extinction patterns prior to and across the KT boundary to be drawn. In the present paper a brief overview is given of these echinoderm faunas, with emphasis on KjiioReBbie cioBa: rio3aHHH Mea, PaHrodi IlajieoreH, echinoids and asteroids. HTJIOKOaCHe, TaKCOHOMHH, CTpaTHTpatpHH, TatJ)OHOMHH.

Kev words: Late Cretaceous, Early Palaeogene, echinoderms. taxon• omy, stratigraphy, taphonomy. Introduction

Résumé The calcareous, multi-element echinoderm skeleton is almost predestined to become fossilised (DONOVAN, A l'exception des échinides. les faunes d'échinodermes de la région- 1991). The often gregarious occurrence of echinoderms type de l'Etage Maastrichtien sont encore plus ou moins terra inco• in many types of marine strata, and the fact that, even in gnito. Ou matériel récemment récolté dans cette région par un groupe de paléontologues professionnels et amateurs comprend de nombreuses the case of dissociated ossicles, material is readily as• et nouvelles pièces qui ont l'avantage supplémentaire d'être bien signed to family, genus or species, makes them an ideal documentées stratimaphiquement. Les collections des musées, et en subject for palaeobiological and palaeoecological studies. particulier celles antérieures à 1975, souffrent généralement du manque de précision en ce qui concerne la position stratigraphique. All species of echinoid, asteroid, ophiuroid and crinoid ( es nom elles collections non seulemenl élargissent noire connaissance from Campanian, Maastrichtian and Danian deposits in des faunes d'échinodermes du Crétacé supérieur (Campanien-Maas- southern Limburg (The Netherlands) and contiguous trichtien) et Paléogène inférieur (Danien) dans la région, mais per• mettent également de tirer des conclusions sur les modelés de diversi• areas in Belgium and Germany are currently being stu• fication et d'extinction de part et d'autre de la limite K/T. Dans cette died (Fig. 1). This is done within the framework of a K/T note, ces faunes d'échinodermes sont brièvement passées en revue en boundary diversification/ extinction project (JAGT, 1998, mettant l'accent sur les échinides et les astéries. 1999a-d; KUTSCHER & JAGT, 1999). This has resulted in Mots-clefs: Crétacé supérieur. Paléogène inférieur, échinodermes, ta• numerous new records. These include not only taxa pre• xinomie, stratigraphie, taphonomie. viously described from elsewhere in northwest Europe, but also quite a lot of new genera and species, especially amongst crinoids, asteroids and ophiuroids. PeiioMC The material consists mostly of dissociated ossicles or portions of skeletons at best (echinoids excepted), but 3a HCKJitoHeitHCM MopcKHx e»eH, (bayHbi HTJIOKO>KHX THUHHHOTO rare finds of well-preserved goniasterid and astropectinid paftoHa MacrpHXTCKoro apyca ocraioTca B ôommeft HJTH asteroids, ophiurid and ophiolepidid ophiuroids, and MeHbinefl creneHH terra incognita. HeoaBHO npe,acTaBJieran,ie bourgueticrinid crinoids are also known. To date, well tpyimoH naneoHTOJioroB, npodpeccHOHajioB H aroÖHTejieft, o6pa3Ubi 3Toro paftoHa enum o&beKTOM MHoroHHCJieHHbix HOBMX over 200 species are recorded from the area. The present

Ha6aK>ÄeHHH, «onojnurrejibHbiM npemiymecTBOM KOTopbix paper provides a brief outline of studies underway as well HBJiaeTca HX cTpaTHrpa<))nHecKa« aoKyMeHTHpoBaHHoert. as a selection of new records from the area. Crinoids are 104 John W. M. JAGT

note is the absence of isocrinids in the Geulhem Member (Early Palaeocene). Comatulids are represented by atelecrinids, pteroco- mids, conometrids, notocrinids and antedomds. and occur throughout the entire Late Cretaceous section. However, in the Campanian and Early Maastrichtian they are com• paratively rare. Their acme is in the Nekum and Meerssen members, where Jaekelometra gr. belgica (JAEKEL, 1902), ./. gr. cancan, (SCHLÜTER, 1878), Semiomclra lenticularis (SCHLÜTER, 1878) and 5. saskiac JAGT, 1999b are very common locally (CBR-Romontbos. ENCI-Maastricht BV and Blom quarries). In addition to centrodorsals representing various ontogenetic stages, such occurrences have also yielded (proximal) brachials, cirrals and pinnules. Brachials with syzygial articulations are fairly common, suggesting these crinoids to have been able to shed arms easily, which in turn would indicate Fig. I Southern Limburg (The Netherlands) and conti• stressful conditions (?increased predation pressure; com• guous areas, showing location of outcrops and quar• pare MISSING. 1997) in shallow-water, subtropical set• ries referred to in the text: tings. At times, species distinction is difficult, particularly 1 - temporary Albertkanaal sections; of Jaekelometra, Amphorometra and Hertha. In Ibis re• 2 - CBR-Romontbos quarry; spect, they resemble extant forms (MESSING, 1997). 3 - CBR-Lixhe quarry; In comparison with the underlying Meerssen Member, 4 - CPL SA quarry; comatulid diversity decreases noticeably in the Geulhem 5 - Ankerpoort-Curfs quarry; Member, with only two forms represented, Hertha gr. 6 - Blom quarry; 7 - Ankerpoort-'t Rooth quarry; mvstica VON HAGENOW, 1840 (?) and Atuatucamctra an- 8 - ENCI-Maastricht BV quarry; nae JAGT, 1999b. 9 - Kunrade; Bourgueticrinids range through the entire section, 10 - Benzenrade; being commonest in the Late Campanian, early Late 11 - Vijlcn. Maastrichtian and Early Palaeocene. Of special note is The inset map of The Netherlands and Belgium the crinoid/ophiuroid lagerstätte at the base of the Grons- shows the area of the main map (shaded). veld Member (ENCI-Maastricht BV quarry; see JAGI et al., 1998). Newly collected slabs which preserve up to ten crinoids, as well as ophiuroids and rare asteroids, demon• strate the impact of storm activity on these crinoid "mea• not illustrated here, and only few figures of ophiuroids are dows". Not only do these allow the density and spatial included, since the chapters describing these echinoderms distribution of "populations" (see BAUMILLER & ROME, have either just come out or are about to be published 1998) to be determined, but also the nature of the sub• (JAGT, 1999b and KUTSCHER & JAGT, 1999, respectively). strate to be analysed in detail. Quite a few crowns have Holothurians are not considered any further; with the penetrated the substrate to depths of almost 10 cm, with exception of dissociated elements of the peristomial ring, arms outspread. ossicles of these echinoderms are extremely rare in the Bourgueticrinids disappear from the section above the area (compare ZELEZNIK, 1985). base of the Emael Member, only to reappear in the Geulhem Member (Albertkanaal sections and Anker• poort-Curfs quarry), with species that are well known Crinoids from Danian strata in Denmark and southern Sweden, namely Bourgueticrinus danicus BRÜNNICH NIELSEN, Known to date from the area are 36 species, in 20 genera, 1913 and Democrinus? maximus BRUNNICH NIELSEN, with all articulate (sub)orders represented. Of these, 3 1915. HÄKANSSON et al. (1996) have recently suggested genera and 6 species are new (JAGT, 1999b). that an important evolutionary phase in the Bourgueticri- Isocrinids (genera Austinocrinus, hocrinusl, Isselicri- nina took place during the earliest Palaeocene, and that nus, Praeisselicrinus! and Nielsenicrinus) are particu• numerous new dorsal cup morphologies arose through larly well represented in the Late Cainpanian (CPL SA neoteny/paedomorphosis. and CBR-Lixhe quarries) and Early Maastrichtian (Vij- Of the infraorder Holopodinidia only a single repre• len/Aaehen area), but do extend into the latest Maastrich• sentative is known, Cvathidium vlieksi JAGT, 1986, which tian. At least one species (possibly two) occur in the is now known from the base of the Vijlcn Member, the shallow-water settings represented by the Nekum and Meerssen Member and the Kunrade limestone facies Meerssen members (Maastricht Formation; Fig. 2), where (Kunrade area). There are no Danian records of this genus they are associated with much commoner comatulids. Of in the study area. Echinoderm faunas near the K/T boundary 105

Finally, roveacrinids have been shown to range to right Fig. 8) allows the ambulacral structure to be described below the K/T boundary, with Birgelenocrinus degraafi in detail, and a direct comparison with the genus JAGT, 1999b and Applinocrinus cretaceus (BATHER, 1924) Plistophvma (see SMITH, 1995; SMITH & JEEEERY, in occurring in the middle/higher Meerssen Member. Of press) to be carried out. the other species, Veugelersia diana JAGT, 1999b, two — the Meerssen Member has yielded diminutive holas¬ stratigraphically highly disjunct occurrences are known; teroids (Pl. 1, Figs. 5-7). These appear to be juveniles one in the early Late Campanian (Benzenrade area) and of the common holasteroid Hemipneustes striatora- one in the Late Maastrichtian (Ankerpoort-'t Rooth quar• diatus (LESKE, 1778), of which literally thousands of ry). adult specimens have been collected in the area. The apparent absence in the area of juveniles of this species has always been a mystery; it may be that Echinoids the fragile tests stood virtually no chance of being preserved in the shallow-water settings represented To date, over 100 species are on record, but diversity is by the Nekum and Meerssen members. still increasing. Notable recent additions and discoveries include the following: JAGT (1998) noted that echinoid faunal composition - the Zeven Wegen Member (CPL SA quarry) has across the K/T boundary varied considerably. Suffer• yielded the highly specialised, bizarre holasteroid ing heavy losses are infaunal selective deposit feeders Hagenowia (PI. 1, Figs. 11, 12), which may prove (hemiasterids), shallow infaunal/semi-infaunal selective conspeciftc with material from Norfolk and northwest deposit feeders (holasteroids), infaunal bulk sediment Germany, and represent a still undescribed member of swallowers (cassidulids), infaunal selective deposit fee• the hlachnorei/elongata lineage. ders (faujasiids) and the epifaunal browser Orthopsis. — the Benzenrade Member near Benzenrade has yielded Epifaunal diversity (cidaroids) is comparable across the the zeuglopleurid Zeuglopleurus rowei GREGORY, boundary, with psychocidarids occurring especially in 1900 (PI. I, Fig. 4), which extends the range of this the Danian. Epifaunal generalists (saleniids) increase in rare species, recorded mostly from white chalk set• diversity across the boundary, and within the Geulhem tings (see SMITH & WRIGHT, 1996), to the early Late Member nearshore hardground grazers (arcopeltids, Campanian. arbaciids) are confined to the upper part. That part has - from the Meerssen Member (ENCI-Maastricht BV also yielded the highest diversity in shallow-water, more quarry) additional spines of the psychocidarid Tylo- protected firm bottom species (phymosomatids, cidar• cidaris inexspectata JAGT & VAN DER HAM, 1995 have oids). been collected, showing it to range to the top of that unit. This species may be a sister taxon of the Early Palaeocene T. oedumi BRUNNICH NIELSEN, 1938, Ophiuroids which in turn may prove to be a junior synonym of T. hardouini (DESOR, 1855). Previous studies of ophiuroids from the study area (e.g.. - A small phymosomatid from the Nekum Member BERRY, 1938) were based almost exclusively on disso• (ENCI-Maastricht BV quarry) preserves spines and ciated ossicles, which generally were poorly preserved. In the lantern, and shows the type of spine referred to as recent years, especially storm-dominated deposits Phvmosoma rutoti LAMBERT, 1898 in the literature, to (Meerssen Member) have yielded many specimens with belong to a genus close to Trochalosoma LAMBERT, discs and (portions of) arms preserved, allowing species 1897. to be better defined. Thus, the pitfalls of combining - of the locally very common Danian saleniid Hvposa- various unrelated types of dissociated ossicles into spe• lenia heliophora (AGASSIZ & DESOR, 1846), a number cies (see discussion in RASMUSSEN, 1950, 1952) can be of specimens with well-preserved lanterns have been evaded. In addition, preliminary observations on preda¬ collected, one of which is illustrated here in PI. 1. tion pressure, based on the number of regenerating arms Figs. 1-3. (see ARONSON, 1987) are possible. - the Meerssen Member (ENCI-Maastricht BV quarry) In view of the fact that most ophiuroid species, parti• has yielded the second, well-preserved specimen of cularly those of the Late Campanian and early Late an apparently new species of the plagiochasmid Maastrichtian, are also known from the Early Maastrich• genus Plagiochasma. MEIJER (1965) noted that it tian of Rugen (NE Germany), KUTSCHER & JAGT (1999) appeared to differ consistently from the Palaeocene decided to base their descriptions mainly on material P. cruciferum (MORTON, 1830) [= P. analis (DESOR, from that locality. Numerous new species, amongst the 1857)]. (sub)families Ophiobyrsinae, Asteronychidae, Euryali- - from the Zeven Wegen Member (CPL SA quarry), dae, Ophiomyxidae, Ophiacanthidae, Ophiuridae, Am- juveniles of the diadematid Centrostephanus? phiuridae, Ophiothricidae, Ophiocomidae, Ophioderma- sp. (PI. 1, Figs. 9, 10) are now known. tidae and Ophiolepididae. Material from the Benzenrade, - new well-preserved material of the stomopneustid Nekum, Meerssen and Geulhem members in the study Winkleria maastrichtensis ENGEL, 1964 (PI. 1, area is complementary. Campanian Maastríchtian Danîan

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lOVf IAI AA «M°R 901 Echinoderm faunas near the K7T boundary 107

In general, the more robust forms such as Ophiomu- — Late Cretaceous species of Crateraster are common siumgranulosum(PI. I, Figs. 16, 17),areoverrepresented (PI. 2, Fig. 22), C. favosus (SPENCER, 1913) occurring in ophiuroid samples. Comparatively more articulated in the Zeven Wegen Member and C. reticidatus remains, such as discs and portions of arms, are known (SCHULZ & WEITSCHAT, 1981) in the Vijlen Mem• of these forms. The Geulhem Member has yielded only ber. Typically Early Palaeocene representatives few species; the most interesting amongst these is a new such as C. anchylus (BRONNICH NIELSEN, 1943) and ophiolepidid (PI. 1. Fig. 13). C. retiformis (SPENCER, 1913) are known from the Geulhem Member, but neither is particularly com• mon. Asteroids — "cryptozonid" (? echinasterid) forms such as the one illustrated by MÜLLER (1953, pi. 10, fig. RAl-2)have Starfishes are the most neglected group amongst the been found associated with ambulacrals and terminal echinoderms of the study area, which may be explained plates in the Zeven Wegen Member (PI. 2, Figs. 1, 2, by the fact that these animals disintegrated rather rapidly 4). upon death and only a jumble of dissociated ossicles — of Metopaster decipiens SPENCER, 1913 (PI. 2, remained (BLAKE, 1989, 1996). However, although more Fig. 3) quite a few associated remains of indivi• or less complete specimens are extremely rare (see e.g., duals have been collected from the Zeven Wegen FAWAS SAINT FOND, 1799; UMBGROVE, 1925), asteroids Member. undoubtedly deserve better. At least 50 species have now — Metopaster undulatus SPENCER, 1913 (PI. 2, Fig. 7), been recognised in strata of Campanian, Maastrichtian which BRETON (1992) placed in his new genus Para- and Danian age. More than half of these have been metopaster, is known exclusively from the Early described previously from elsewhere in Europe, in parti• Maastrichtian portion of the Vijlen Member; a com• cular from white chalk facies types. These include the parable (new?) form, with a pronounced ornament of following: pits of varying size (PI. 2, Fig. 10), occurs in the - representatives of the studlandensis-alseni-peakei Zeven Wegen Member. lineage of Nymphaster, which have been shown to - the enigmatic goniasterid Chomataster acules SPEN• constitute good index fossils (GALE, 1987b, 1989; CER, 1913, ranging from the early Late Campanian to BRETON, 1992) in the early and late Late Campanian the Middle Danian (Late Danian in Denmark; RAS• (PI. 2, Figs. 8, 19, 20). MUSSEN, 1945), of which a fairly well-preserved in• - Nymphaster spenceri (RASMUSSEN, 1950) and N. dividual is known from the Geulhem Member of the wrighti (RASMUSSEN, 1950), well known from the temporary Albertkanaal sections (Vroenhoven- Maastrichtian of northern Germany and Denmark. Riemst). - Lophidiasterpygmaeus SPENCER, 1913 (PI. 2, Fig. 9), — Ophryaster oligoplax (SLADEN, 1891) (PI. 2, Figs. 5, which first occurs in the Zeven Wegen Member, and 6), of which several well-preserved arm fragments is a minor constituent of asteroid faunas in the re• and numerous dissociated marginals, some preserving mainder of the Gulpen Formation and the lower part granules, have been collected from the Zeven Wegen of the Maastricht Formation. A find of associated Member; in the Vijlen Member, O. magnus SPENCER. ossicles of a single individual suggests that MULLER'S 1913 occurs in places. (1956) description of this taxon is in need of a revi• sion. Typically Early Palaeocene species such as Metopaster - the sphaerasterid Valettaster (PI. 2, Fig. 15), of which spencerii BRÜNNICH NIELSEN, 1943 (PI. 2, Fig. 17), M. at least two species occur, one of them possibly new kagstrupensis BRÜNNICH NIELSEN, 1943 and Astropecten (Zeven Wegen Member), the other assignable to V. punctatus (BRÜNNICH NIELSEN, 1943) (PI. 2, Fig. 16) are ocellatus (FORBES, 1848), a long-ranging taxon (see known from the Geulhem Member, and it is in part on BRETON, 1985). these forms that RASMUSSEN (1965) based his correlation — Metopaster tumidus SPENCER, 1913, of which typical of that unit with the lower part of the Danish Bryozoakalk examples are known only from the Early Maastrich• (Stevns Klint). tian portion of the Vijlen Member (PI. 2, Fig. 21); forms assignable to SCHULZ & WEITSCHAT'S (1975) In addition to the above-mentioned taxa, which allow M. praetumidus occur in the Zeven Wegen Mem• interregional correlations with localities elsewhere in ber. northwest Europe, the asteroid faunas in the study area also contain what appear to be endemic elements, although use of the term '-endemic" should in fact be avoided. It may well be that the "typical Maas• tricht tuffaceous chalk" facies was distributed much more extensively over northwest Europe, but that these Fig. 2 — Lithostratigraphy and biozonation of Campanian- strata were eroded completely by subsequent transgres- Maastrichlian strata in the type area of the Maas• sive phases. This, in fact, goes for all echinoderm groups, trichtian Stage (from JAGT, 1999a). not only for starfish. New finds include: 108 John W. M. JAGT

Table 1 — Stratigraphie distribution (by lithostratigraphic member; see Fig. 2) of echinoderm taxa from the extended type area of the Maastrichtian Stage, known to date. Abbreviations are as follows: VF - Vaals Formation, ZW - Zeven Wegen Member. B - Bcutenaken Member, Vij - Vijlen Member, Li - Lixhe 1-3 members, L - Lanayc Member, Va - Valkenburg Member, Gr - Gronsveld Member, S - Schiepersberg Member, E - Emael Member, N - Nekum Member, M - Meerssen Member, Ge - Geulhem Member.

VF ZW B Vij Li L Va Gr S E N M Ge Echinoids Tylocidaris bruennichi X Ty. hardouini X Ty. inexspectata X Temnocidaris nigelliensis X X T. dánica X T. serrata X T. uhaghsi X Ctenocidarisl distincta X Cidarisl forchhammeri X C? rosenkrantzi X Goniocidaris: sp. X Phyllacanthus sp. X echinothuriid X X X Centrostephanusl sp. X X X X X Orthopsis miliaris X X pedinoid X Salenia anthophora X S. bélgica X

S. bonissenti \ X X S. gr. nutrix X S. sigillata X X S. sp. nov. X S. heberti X S. maestrichtensis X X X Salenocidaris mínima X Sal. obnupta X Hyposalenia helíophora X Codiopsis disculus X X Goniopygus minar X G. heberti X G. sp. nov. X Phymosoma granulosum X X X Circopeltis maastrichtensis X C. sp. X Phymotaxis tournoueri X Thylechinus sp. nov. X Gauthieria grossouvrei X G. mosae X G. pseudoradiata x X X X X X X X X G. maeandrina Gauthiosoma princeps X X Echinoderm faunas near the K/T boundary 109

VF ZW B Vij Li L Va Gr s E N M Ge Ga. krimica X Trochalosomal rutoti X X X X X X X Mierapsidia salís X X Micropsia sp. nov. X Winkleria maastrichtensis X X Zeuglopleurus rowei X Coenholectypus macrostomus X X Galerita stadensis X X G.l hemisphaericus X GP. sulcatoradiatus X Adelopneustes montainvillensis X A. boehmi X Echinogalerus bélgicas E. muelleri E. pusillus X E. transversas X X E.l vetschauensis Nucleopygus coravium X X X X X N. scrobiculatus X X X X X X Gitolampas oblongas X Procassidulus lapiscancri X X X P. elongatus X Rhynchopygus marmini X X Rhyncholampas macari X X Faujasia apicalis X X X Catopygus fenestra tus X X X X X X Oolopygus pyriformis X X X X X X Plagiochasma cruciferum X P. sp. nov. X Hagenowia sp. X Cardiaster granulosus X X X X X X X C. rutoti X X X Cardiotaxis heberti X Echinocorys gr. cónica X E. gr. conoidea X E. gr. gibba X E. gr. humilis X E. gr. limburgica X E. gr. ovata X E. gr. subglobosa X E. gr. pyramidata X X Galeota papulosa basiplana X Hemipneustes oculatus X X X H. striatoradiatus X X X X X X Micraster gr. schroederi/glyphus X X M. stolleyi X Cyclaster platornatus X 110 John W. M. JAGT

VF ZW B Vij 1.1 1. Va Gr s E N M Ge Diplodetus spp. X X D. cf. americanus X D. parvistella X X X D. duponti X X D. bucardium X X Hemiaster gr. aquisgranensis X X X X X X X H. prunella X X X X H. koninckanus X X X X Leymeriaster eluvialis X X X L. maestrichtensis X X L. sp. nov. X Linthial breviuscula X LiP. sp. X Paraster sindensis X Ophiuroids Ophiosmilaxl sp. nov. X X Asteronyxl sp. nov. X

Trichasterl omatus X X X X X X X \ T.l sp. X X X X Ophiomyxal sp. nov. X X Ophiomyxal jekerica X X X Ophioscolexl sp. nov. 1 X Ophioscolexl sp. nov. 2 X X Ophiacanthal dánica X X X X X Ophiacanlluí! sp. nov. 1 X Ophiacanthal sp. nov. 2 X X Stegophiural hagenowi X X X SP sp. nov. X Ophioctenl sp. nov. X Felderophiura vanderhami X X X Ophioplinthaca? fuerstenbergii X X Amphiura"! sp. nov. X X amphiurid gen. et sp. nov. 1 X amphiurid gen. et sp. nov. 2 X Ophiothrixl sp. nov. 1 X X X X Ophiothrixl sp. nov. 2 X X Ophiactisl sp. nov. X Ophiocomal senonensis X X X X Ophioderma? sp. nov. X X Ophiodermal substriatum X X X Ophiarachnal sp. nov. X Ophiotitanos serrata X X X X X X X X Ophiolepisl sp. nov. 1 X Ophiolepis? sp. nov. 2 X X Ophiolepisl sp. nov. 3 Ophiomusium sp. nov. 1 X Ophiomusium granulosum X X X X X X X X Echinoderm faunas near the K/T boundary 111

VF ZW B Vij Li L Va (ir S E N M Ge Ophiomusium sp. nov. 2 X Ophiomusium sp. nov. 3 X ophiolepidid gen. et sp. nov. X Crinoids Austinocrinus bicoronatus X lsocrinu/l sp. X 1.1 lanceolatus X Isselicrinus buchii X Praeisselicrinusl limburgicus X Nielsenicrinus carinatus X N. agassizii X N. ewaldi X X Jaekelometra gr. bélgica X X

J. gr. cóncava X X J.l defectiva X X Placometra gr. laticirra X X X X

Atuatucametra annae X Amphorometra gr. conoidea X X X Semiometra impresso X

S. lenticularis X X

S. saskiae X X Loriolometra retzii X Hertha gr. plana X X

H. gr. pygntea X

H. gr. mystica X Dunnicrinus aequalis X X X X Bourgueticrinus aff. baculatus X B. sp. X X

B. bruennichinielseni X B. aff. brydonei X X B. constrictus X X

B. danicus X B. hureae X

B.l suedicus X

Democrinus? maximus X -Monachocrimts gal Ileus" X

Cyathidium vlieksi X X

Applinocrinus cretaceus X X X X X X X Birgelenocrinus degraafi X

Veugelersia diana X X Asteroids astropectinid sp. X

Astropectenl punctatus X

A.'? sp. nov. X

Aldebarania sp. nov. X

Lophidiaster pygmaeus X X X X X X X

Coulonia sp. nov. X X 112 John W. M. JAGT

VF ZW B Vij Li L Va Gr S E N M Ge benthopectinid X X X Stauranderaster miliaris X Aspidaster sp. X Pvcinaster crassus X X P. danicus X Valettaster granulatus X V. ocellatus X X V. sp. (nov.?) X Metopaster sp. nov. 1 X M. spencerii X M. kagstrupensis X M. sp. nov. 2 X M. sp. nov. 3 X M. sp. nov. 4 X M. decipiens X M. sp. nov. 5 X M. sp. nov. 6 X M. praetumidus X M. tumidus X M. undidatus X Ceramasterl sp. X Recurvaster hlackmoreil X R. radiatus X R. sp. nov. 1 X X R. sp. nov. 2 X Chomastaster acules X X X X X X X X X goniasterid sp. 1 X X X X goniasterid sp. 2 X X X Ophryaster oligoplax X 0. magnus X OP maastrichtensis X Comptoniaster sp. nov. X Crateraster favosus X C. reticulums X C. sp. nov. X C. anchylus X C. retiformis X Nymphaster studlandensis X N. alseni X N. spenceri X N. wrighti X NP sp. X asteriid(s) X X "cryptozonids" X X Echinoderm faunas near the K/T boundary 113

- apparently juvenile goniasterids, closely related to 1997), VILLIF.R (1996) and VILLIER et al. (1997). One Crateraster (PI. 1, Fig. 14), from the Meerssen Mem• of the new species, Metopaster sp. nov. (PI. 2, Fig. 18) ber (Blom quarry). is reminiscent of M. calcar SPENCER, 1913, from the many paxillosidans, such as possible radiasterids Santonian-Early Campanian of southern Sweden (see (PI. 2, Fig. 13) and astropectinids, e.g. the form illu• GALE, 1987a, pi. 8, figs. 12-21) and of M. brom/evi strated in PI. 2, Fig. 12, which RASMUSSEN (1965, pi. 8, GALE, 1987a (pi. 2, figs. 14-16; pi. 3, figs. 1-5) from fig. 13) referred to as Asteropecten n. sp. aff. cottes- the late Early Campanian of the same area. woldia [sic]. Cainozoic astropectinids (see e.g., RAS• - remains of at least two individuals of what appears to MUSSEN, 1972; KACZ.MARSKA, 1987) are in need of a be a new species of Recurvaster, seemingly closely modem revision; only rarely have species been based related to and a possible precursor of the Early Pa• on such well-preserved remains as those recently laeocene R. mammillatus (GABB, 1876), are known described by NOSOWSKA (1997). Forms closely re• from the uppermost Meerssen Member (PI. 2, Figs. 23, lated to or assignable to the genera Tethvaster (see 24) of Blom quarry. e.g., HALL & MOORE, 1990; BRETON, 1995), Dipsa- - dissociated ossicles of asteriids (PI. 1, Figs. 15, 19), caster (see BRETON el al., 1995), Coidonia (= Cunea- closely comparable to material from the Cenomanian- ster; see HESS & BLAKE, 1995) and the otherwise Coniacian of France as illustrated by BRETON & exclusively North American Aldebarania (see BLAKE FERRE (1995). Asteriids have rarely been recorded & STURGEON, 1995) are known from the Zeven We• as fossils, and for that reason it comes as no surprise gen Member and the higher Maastricht Formation that new finds almost invariably represent new genera (Emael, Nekum and Meerssen members; PI. 2, and/or species (see e.g., BLAKE, 1990a; BLAKE & Fig. 11), in particular. From flint nodules in the upper PETERSON, 1993; BLAKE et al., 1996; BLAKE & AR- Nekum Member (CBR-Romontbos quarry), three ONSON, 1998). well-preserved individuals are known of an astropectinid referable to Aldebarania (D.B. Blake, pers. Reference is made to JAGT (1999d) for more details and comm.). for a discussion of functional morphology and palaeoe- - stauranderasterids, especially from the Geulhem cology of these asteroid faunas, based on literature data Member, appear close to Stauranderaster miliaris (BLAKE, 1989, 1990b). BRÜNNICH NIELSEN, 1943 (PI. 2, Fig. 14); the Meers• sen Member has yielded another form (PI. 2, Fig. 25). Acknowledgements rare benthopectinids (PI. 1, Fig. 18), comparable to

material illustrated by BLAKE (1973, 1984). 1 wish to thank the following colleagues for assistance in various - various new species of the goniasterid Metopaster, ways: M.J. Van Birgelf.n, D.B. Blake. G.J. Boekschoten, M.J.M. this genus in particular appears to have been very Deckers, S.K. Donovan, R.W. Dortangs, A.S. Gale. R.W.J.M. successful in establishing local species, as offshoots Van di k Ham, H. Hess. J. INDEHERBERGE, Saskia M. Kars. P. Van Knippenbero. M. Kutscher, M.M.M. Kuypers, J.P.H. Reynders of the parkinsoni lineage. GALE (1987a) erected a and J. Snellings. The managements of CPL SA-Haccourt, CBR- number of such short-lived offshoots, which are par• Lixhe, ENCI-Maastricht BV. Blom and Ankerpoort-Curfs allowed ticularly typical of marginal settings, e.g., coarse• access to their quarries, which is much appreciated. This research was supported (in part) by the Geosciences Foundation (GOA) with grained biocalcarenites in southern Sweden. Compar• financial aid from the Netherlands Organisation for Scientific Research able forms have been discussed by BRETON (1992, (NWO).

References

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Patterns and processes of heterochrony in recovery subsequent to the Cretaceous-Tcrtiarv boundary - the Mesozoic goniasterid sea-stars, l.ethaia. 30: 135-144. European example. Abstracts. Sixth North American Paleonto- BRETON, G., BILOTTE, M. & SKIRO, G., 1995. Dipsacaster jadeti logical Convention, Washington DC, 1996: I pp. sp. nov., Astropectinidae (Asteroidea, Echinodennata) du HALL, R.L. & MOORE, S., 1990. ITethvaster albertensis, a Maastrichtien des Petites Pyrénées (France). Bulletin trimestriel Late Cretaceous (Turanian) sea star from the Cardium For• de la Société géologique de Normandie et des Amis du Muséum mation, Alberta, Canada. Journal of Paleontology, 64: 1045¬ du Havre, 82: 35-42. 1049. BRI ION, G. & FERRÉ, B.. 1995. Première observation d'élé• HESS, H. & BLAKE, D.B., 1995. Coulonia platyspina n. sp. . a ments squelettiques d'Asteriidae (Asteroidea, Echinodennata) new astropectinid sea star from the Lower Cretaceous of Mor• dans les craies du Cénomanien au Coniacien du Bassin de Paris occo. Eclogue geologicae Helvetiae, 88: 777-788. (fiance). Revue de Micropaléontologie, 38: 299-309. JAEKEL, O., 1902. Uber verschiedene Wege phylogenetischer BRÜNNICH NIELSEN. K.. 1913. Crinoidernc i Danmarks Kridtaf- Entwicklung. Verhandlungen des funfien intemationalen zool¬ Icjringer. Danmarks geologiske Undersôgelse, (2)26: 1-120. ogischen Congress, Berlin. 1901: 1058-1117. BRÜNNICH NIELSEN, K., 1915. Rhizocrinus máximas n. sp. og JAGT, J.W.M., 1986. Cvathidiuin vlieksi, a new holopodid cri- nogle Bemœrkninger om Bourguetierinus danicus Br. N. Med- noid from the Upper Maastrichtian (Late Cretaceous) of south• delelserfra dansk geologiske Forening, 4: 391-394. ern Limburg, The Netherlands. Geologie en Mijnbouw, 65: 215¬ BRÜNNICH NIELSEN, K.. 1938. Faunaen i /Eldre Danium ved 221. fCorporalskroen. Meddelelser Ira dansk geologiske Forening, 9 JAGT, J.W.M., 1998. Late Cretaceous-Early Palaeogene echi- (1937): 118-126. noderm faunas and the K/T boundary in the Maastrichtian type BRÜNNICH NIELSEN, K., 1943. The asteroids of the Senonian and area. In: KOPAEVICH, L.F. & DHONDT, A.V. (org.). INTAS Danian deposits of Denmark. Dct kongelige Danske I idenska- Project 1994-1414, Bio-events at the K/T boundary on the bernes Selskah. Biologiske Shifter, (2)5: 1-68. southern margin of the white chalk sea - paleobiology, palaeo- DESOR, E., 1855-1858. Synopsis des Échinides fossiles. biogeography, sequence stratigraphy, geochemistry and geo- Echinoderm faunas near the K/T boundary 115 chronology. Final Meeting, Moscow March 23-25, 1998: 17, MÜLLER, A.H., 1956. Zur genaueren Kenntnis von Lophidiaster 18. pvgmaeus (Asterozoa) aus der Schreibkreide (Maastricht) von JAGT, J.W.M., 1999a. Late Cretaceous-Early Palaeogene echi- Rügen. Geologie, 5: 642-651. noderms and the K/T boundary in the southeast Netherlands and NOSOWSKA, E., 1997. Asteroids from the Nawodzice Sands northeast Belgium - Part 1: Introduction and stratigraphy. (Middle Miocene; Holy Cross Mountains, Central Poland). Scripta Geológica, 116A: 1-57. Acta geológica polonica, 47: 225-241. JAGT, J.W.M., 1999b. Late Cretaceous-Early Palaeogene echi- RASMUSSEN, H.W., 1945. Observations on the asteroid fauna of noderms and the K/T boundary in the southeast Netherlands and the Danian. Meddelelser fra danske geologiske Forening, 10 northeast Belgium - Part 2: Crinoids. Scripta Geológica, 116B: (1944): 417-426. 59-256, 46 pis. RASMUSSEN, H.W., 1950. Cretaceous Asteroidea and Ophiur- JAGT, J.W.M., 1999c. Late Cretaceous-Early Palaeogene echi- oidea with special reference to the species found in Denmark. noderms and the K/T boundary in the southeast Netherlands and Danmarks geologiske Undersgelse, (2) 77: 1-134. northeast Belgium - Part 4: Echinoids. Scripta Geológica, 122 RASMUSSEN. H.W., 1952. Cretaceous Ophiuroidea from Ger• (in press). many, Sweden, Spain and New Jersey. Meddelelser fra dansk JAGT, J.W.M., 1999d. Late Cretaceous-Early Palaeogene echi- geologiske Forening, 12 (1951 ): 47-57. noderms and the K/T boundary in the southeast Netherlands and RASMUSSEN, H.W., 1965. The Danian affinities of the Tuffeau northeast Belgium - Part 5: Asteroids. Scripta Geológica, 123 de Ciply in Belgium and the "Post-Maastrichtian" in the (in press). Netherlands. Mededelingen van de Geologische Stichting. JAGT, J.W.M., DONOVAN, S.K., DECKERS, M.J.M., DORTANGS, n.s., 17: 33-40. R.W., KUYPERS, M.M.M. & VELTKAMP, C.J., 1998. The Late RASMUSSEN, H.W., 1972. Lower Tertiary Crinoidea, Asteroidea Maastrichtian bourgueticrinid crinoid Dunnicrinus aequalis and Ophiuroidea from northern Europe and Greenland. Bio- (d'Orbigny, 1841) from The Netherlands and Belgium. Bulletin logiske Shifter fra det danske Videnskabeliges Selskab, 19: de I Institut royal des Sciences naturelles de Belgique, Sciences 1-83. dela Terre, 68: 129-154. SCHLÜTER, C, 1878. Ueber einige astylide Crinoiden. Zeit• JAGT, J.W.M. & VAN DER HAM, R.W.J.M., 1995. First record of schrift der deutschen geologischem Gesellschaft, 30: 28-66. the echinoid genus Tvlocidaris Pomel 1883 from the type SCHULZ, M.-G. & WEITSCHAT, W., 1975. Phylogenie und Maastrichtian of the Netherlands. Paläontologische Zeitschrift. Stratigraphie der Asteroideen der nordwestdeutschen Schreib• 69: 233-239. kreide. Teil I: Metopaster/Recurvaster- und Callider- KACZMARSKA, G., 1987. Asteroids from the Korytnica Basin ma/Chomataster-Gruppe. Mitteilungen aus dem geolo• (Middle Miocene; Holy Cross Mountains, Central Poland). gisch-paläontologischen Institut der Universität Hamburg, Acta geológica polonica, 37:131-144. 44: 249-284. KUTSCHER, M. & JAGT, J.W.M., 1999. Early Maastrichtian SCHULZ. M.-G. & WEITSCHAT, W., 1981. Phylogenie und Stra• ophiuroids from Rügen (northeast Germany) and Mon (Den• tigraphie der Asteroideen der nordwestdeutschen Schreibk• mark). In: JAGT, J.W.M. Late Cretaceous-Early Palaeogene reide. Teil II: Crateraster/Teichaster-Gmppe und Gattung echinoderms and the K/T boundary in the southeast Nether• Ophryaster. Mitteilungen aus dem geologisch-paläontolo• lands and northeast Belgium - Part 3: Ophiuroids. Scripta gischen Institut der Universität Hamburg, 51: 27-42. Geológica, 121 (in press). SLADEN, W.P., 1891-1893. A monograph of British fossil Echi• LAMBERT, J., 1897. Note sur quelques Echinides éocènes de nodermata from the Cretaceous formations, 2. The Asteroidea l'Aude, I. Endocycles. Bulletin de la Sociêté géologique de and Ophiuroidea, 1-2. Monographs of the Palaeontographical France, (3) 25: 483-517. Society of London, 1891-1893: ii + 1-66. LAMBERT, J., 1898. Note sur les Échinides de la craie de Ciply. SMITH, A.B., 1995. 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MESSING, CG., 1997. Living comatulids. in: WATERS, J.A. & SPENCER, W.K., 1913. The evolution of the Cretaceous Aster• MAPLES, CG. (Eds), Geobiology of echinoderms. Paleontoló• oidea. Philosophical Transactions of the Royal Society of Lon• gica! Society Papers, 3: 3-30. don, B204: 99-177. MORTON, S.G., 1830. Synopsis of the organic remains of the UMBGROVE, J.H.F., 1925. Asteroidea uit het Maastrichtsche ferruginous sand formation of the United States, with geologi• Tufkrijt. Verhandelingen van het Geologisch-Mijnhouwkundig cal remarks. American Journal of Science, 17: 274-295. Genootschap voor Nederland en Kolonien, geol. serie, 7: 207¬ MÜLLER, A.H., 1953. Die isolierten Skelettelemente der Aster• 212. oidea (Asterozoa) aus der obersenonen Schreibkreide von Rü• VILLIER, L., 1996. Les Goniasteridae (Asteroidea, Echinoder• gen. Geologie (Beiheft), 8: 3-66. mata) du Campanien d'Aquitaine; ontogenèse, évolution et 116 John W. M. JAGT paléoécologie. Université de Bourgogne, Centre des Sciences Université de Liège, Faculté des Sciences, Laboratoires de de la Terre, 50 pp. (unpublished thesis). Paléontologie, 104 pp. (unpublished thesis).

ViLLiER, L., BRETON, G. & NÉRAUDEAU, D., 1997. Contexte paléoécologique, biodiversité et signification biostratigraphi- John W. M. JAGT que des asteridés dans le Campanien stratotypique. Annales Natuurhistorisch Museum Maastricht de la Sociale Géologique du Nord, (2) 5: 181-188. P.O. Box 882 VON MACHNOW, F., 1840. Monographie der Rügen'schen Krei- NL-6200 AW Maastricht deversteinenmgen. Abtheilung II. Radiarien und Annulaten, The Netherlands nebst Nachträgen zur I. Abtheilung. Neues Jahrbuch für Miner• [mail(«!nhmmaastricht.nl] alogie. Geognosie, Geologie undPetrefactenkunde, 1840: 631¬ 672. ZELEZNIK, A„ 1985. Analyse descriptive et quantitative des meso-et microfossiles du contact Campanien-Maastrichtien Typescript submitted: December I, 1998. dans le Nord-Est de la Belgique et le Limbourg néerlandais. Revised typescript received: February 2, 1999.

PLATE 1

Note — Data on provenance of material in Plates 1 and 2, as well as repository and registration numbers of specimens illustrated, are supplied by JAGT (1999b-d) and KUTSCHER & JAGT (1999), to which reference is made.

Figs. 1-3 - Hyposalenia hetiophora, apical and lateral views of test (x 6), and associated lantern (x 23). Fig. 4 — Zeuglopleurus rowei, apical view, x 9. Figs. 5-7 — Hemipneustes striatoradiatus juv., apical, oral and lateral views, x 9. Fig. 8 — Winkleria maastrichtensis, oblique lateral view, x 17.5. Figs. 9, 10 — Centrostephanusl sp. juv., lateral and oblique apical views, x 15. Figs. 11,12 — Hagenowia sp. (?nov.), rostra, x 15 and x 12, respectively. Fig. 13 — Ophiolepididae n. sp. , lateral arm plate, x 30. Fig. 14 — Goniasteridae indet. juv., x 6.5. Figs. 15, 19 — asteriid indet.. x 20 and x 26, respectively. figs. 16, 17 — Ophiomusium granulosum, proximal arm portions, x 16 and x 5, respectively. Fig. 18 — benthopectinid indet., x 21.

Pi \II 2

Figs. 1,2,4 indeterminate '•cryptozonid" (? echinasterid), marginals, ambulacrals and terminal plate; x 13 (11. \ 19(2) and x 25 (4). Fig. 3 Meiopaster decipiens, ultimate superomarginal, x 9. Figs. 5, 6 Ophryaster oligopla.x, marginals preserving granules, x 10. Fig. 7 Meiopaster undulutus, ultimate superomarginal, x 7.5. Fig. 8 Nymphaster studlandensis, distal marginal, x 17. Fig. 9 Lophidiaster pygmaeus, superomarginal, lateral view, x 20. Fig. 10 "Parametopaster" (sp. nov.?), oblique view of ultimate superomarginal, x 8. Fig. 11 astropectinid indet., superomarginal, x 18. Fig. 12 astropectinid (sp. nov.?), inferomarginal. x 18. Fig. 13 radtasteridf?) indet., superomarginal, x 20. Fig. 14 stauranderasterid. x 11. Fig. 15 Valettaster sp. (nov.?), x 7. Fig. 16 Astropecten punctatus, interradial superomarginal. x 18. Fig. 17 Meiopaster spencerii, median superomarginal, x 4.5. Fig. 18 Meiopaster sp. nov., ultimate superomarginal, x 4. Figs. 19, 20 Nymphaster alseni, interradial superomarginal, x 3. Fig. 21 Meiopaster tumidus, ultimate superomarginal, x 3. Fig. 22 Crateraster reticulatus, median supero- and inferomarginal, x 3. Figs. 23, 24 Recurvaster sp. nov., median superomarginal, x 3. Fig. 25 stauranderasterid, x 3.5.

118 John W. M. JAGT

PLATE 2